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NOT TO BE TAKEN FROM THIS ROOM 


SEASONAL DRIFT IN CARBOHYDRATE CONCEN¬ 
TRATIONS IN GROWING BARLEY 


J. Kastelic 

Department of Flant Science 


University of Alberta 





















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SEASONAL DRIFT IN CARBOHYDRATE CONCEN¬ 
TRATIONS IN GROWING BARLEY 


J. Kastelic 

Department of Plant Science 


A THESIS 

submitted to the University of Alberta 
in partial fulfilment of the 
requirements for the degree of 
MASTER OF SCIENCE 


Edmonton, Alberta 
April, 1945 










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TABLE OF CONTENTS 


• \ 


Page 


Introduction . 1 

Review of Literature . 3 

Materials and Methods.«. IS 

Materials . IS 

Method of collection .. 13 

Preparation of material . 14 

Method of analysis ...* 17 

Technique used in estimation of sugars S3 

Experimental Results . S4 

Seasonal drift in sugar and dry matter 

content . S4 

Discussion and Summary . 61 

Acknowledgements .. 67 

References . 68 



















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SEASONAL DRIFT IN CARBOHYDRATE CONCEN¬ 


TRATIONS IN GROWING BARLEY 


J. Kastelic 


INTRODUCTION 


Since the potential supply of raw fructose in the 
form of inulins and fructosans is now known to he enormous, 
attention has been directed towards possibilities of commer¬ 
cial utilization. If fructose were to become available in 
large quantities at the same price as sucrose, its distinctive 
properties should earn for it a permanent place on the carbo¬ 
hydrate market. It is the sweetest and most soluble of all 
sugars. Its presence in syrups prevents the crystallization 
of the other sugars present and, in contrast to sucrose which 
is subject to inversion change, its flavor is permanent. It 
is also in demand as an ingredient of medicinal preparations. 

Economic and technological barriers have prevented 
the development of a fructose industry in the past. There is, 
however, reason to believe that the technological difficulties 
have been largely overcome. At the Iowa State College and at 
the University of Illinois processes for manufacturing fructose 





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from the tubers of Jerusalem Artichokes have been in success¬ 
ful pilot scale operation for a number of years. 

It should be emphasized that the economic feasibility 
of a fructose industry in Canada must be left in doubt until 
sufficient over-all data on yield and a comprehensive survey 
of sources of raw fructosans become available. It may also 
be pointed out that the effect of soil and weather conditions 
on the fructosan accumulation in plants is at present not too 
well understood. While the occurrence of fructosans and their 
importance in the general carbohydrate metabolism in plants 
have received a great deal of attention from a number of 
biochemists at the Royal Institute of Technology in G-reat 
Britain, no attempt has been made to investigate commercial 
possibilities of using fructosans as raw material for a fruc¬ 
tose industry. From the conclusions reached by fructosan 
chemists, it would appear, on the surface at least, that the 
accumulation of fructosans in plants is the result of many and 
probably complex factors. 

Nevertheless, certain analytical results obtained 
in the biochemical study of rye, wheat, and barley indicate 
some possibility of finding cereal and forage crops suitable 
for commercial production of fructose. Norman (13) in Great 
Britain reported in 1936 that the fructosan constituted 34 fo 
of the oven dry weight of the above-ground portions of rye 
grass after about six weeks of growth. Since fructose aris¬ 
ing from sucrose was included in his estimate this value 




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would appear to be much, too high. This has been shown in 
subsequent investigations by others (4). Notwithstanding the 
failure to confirm Norman*s results there remains some pos¬ 
sibility of finding relatively high concentrations of fruc- 
tosans in certain plants grown under conditions which promote 
the storage of this sugar. 

The investigation contained in this report was 
undertaken as a preliminary survey of the effect of some of 
the factors which influence the fructose and fructosan con¬ 
tent of barley grown under field conditions. 

REVIEW OF LITERATURE 


A detailed review of the fructosans in monocoty¬ 
ledons can be obtained by referring to Archbold (4). It is 
stated that the earliest study on this compound was made by 

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Kuhnemann (1875), Munstz (1878-1886), and Ludwig and Muller 
(1873). The discovery of fructosan in cereals resulted from 
the attempts of agricultural chemists to isolate dextrins as 
intermediate products between sugar and starch during the 
development of the grain. So much controversy arose at this 
time as to the presence or absence of dextrins in monocoty¬ 
ledons that little if any significance was given to the 
discovery of the fructose polymer. While a number of attempts 
were made to prepare a pure fructosan from plant extracts, the 



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- 4 - 


methods in use at that time led to isolation of more or less 
impure substances. This resulted in a variety of descriptions 
of the compound so that even to this day the nomenclature of 
these carbohydrates is far from satisfactory. 

The more modern work, especially that by Augem (1938), 
Belval (1934, 1933, 1939), Colin and Chaudin (1933), Challinor 
et al (4); Norman (1936) (14); Archbold (1938) (3); and Bar- 
nell (1938) (10), has confirmed the presence of fructosans in 
the plants examined by the early workers. It is now well 
established that the fructosans are usually found as one of 
the soluble sugars in the cereals and forage crop plants, and 
that they may form an appreciable part of the total sugar at 
certain stages during growth. 

There still remains a great deal of research to be 
done on the chemistry of the fructosan molecules, but it is 
now generally accepted that they yield only fructose on acid 
or invertase hydrolysis and that their chemical constitution 
may differ slightly according to the source. Full details of 
their chemistry properly belong to the field of pure chemistry 
and may be obtained from the original papers of Hibbert et al 
and Challinore et al (4), Haworth et al (13), and Reich, W.S. 
(15). 

It has been only in relatively recent years that 
satisfactory methods for the quantitative estimations of 
fructosans have been developed and almost without exception 
all of the work has been done by investigators at the Research 



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Institute of Plant Physiology, Imperial College of Science 
and Technology, London. The earlier work was done in France 
and Germany, and in the literature reviewed no reference to 
fructosan is reported in any of the work on plant sugars 
carried out on this continent* 

From the available information it can be stated 
that the amounts of fructosan found in the several plants 
studied vary within wide limits and are subject to the same 
marked seasonal changes and environmental conditions that 
affect the other sugars. In a series of investigations by 
Colin and Belval (4) on the changes in sugar concentrations in 
stems and heads of cereals it was observed that, as the spike 
emerged, fructosans began to accumulate in the stems and the 
concentration reached a maximum when full growth of the plant 
was reached. In wheat stems the value reached was 5.08$ and 
at the onset of head formation the heads contained 6 to 8$ of 
the fresh weight. No fructosans were found in the leaves. 

Archbold^ and Barter f s (6) successful isolation of 
fructosans from barley leaves reopened the question of the 
significance of fructosans in carbohydrate metabolism and 
marked the real beginning of detailed study of the role they 
play. By 1940 Archbold (4) had firmly established that fruc¬ 
tosans may be distributed throughout the whole plant. 

Earlier workers, notably Colin and Belval (4) 
supported the view that stored sugars, including fructosans, 
were stored during the early stages of the plant*s growth and 


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that these reserve stores of sugar were drawn upon when the 
heads began to develop♦ Barnell (9) (10), working with wheat 
and Archbold (3) (5), working with barley, confirmed Colin and 
Belval’s (4) results on the seasonal drift of sugar concentra¬ 
tions. When, however, their results were examined on the 
basis of quantity per plant, it became clearly evident that 
the stored sugars were not available in the sense that they 
can be used as substrates for further growth at a site distant 
from that at which they originally accumulate. Archbold (3) 
(5), Archbold and Mukerjee (8), and Archbold and Datta (7) 
observed that storage of sugar and fructosan was simply a 
balance of supply over demand and that the sudden onset of the 
large loss of these sugars from the plant tissues associated 
with the development of the heads is unrelated. The head is 
therefore entirely dependent on primary assimilate translocated 
to it or produced in situ and immediately and irreversibly 
synthesized. 

A consideration of results of the investigations of 
Archbold et al leads to the conclusion that only conditions 
which promote the production of primary assimilates by green 
parts of the plant in excess of immediate growth requirements 
will result in storage of sugars. It is also to be noted 
that conditions which promote sugar storage result also in 
the production of relatively large amounts of fructosan. 

Losses from the plant after the maximum is reached are 
related to levels of concentrations reached. 



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It has been observed by Archbold (3) and Russell 
(17) that mineral deficiencies have rather marked effect on 
the amounts of sugars stored* An indication of what was ob¬ 
served when barley was grown under conditions of mineral 
deficiencies (taken from Archbold’s and Russell’s work) 
appears in the following tables: 


TABLE A 

Total sugar and fructosan content of barley 
plants grown in sand and receiving 
nitrogen-deficient dressings 




% fresh weight 




Leaves 

Stems 

Heads 



Control -N 

Control -N 

Control 

-N 

Total sugar 

1.58 3.04 

2.03 5.76 

4.79 

4.81 

Fructosan 

0.13 1.15 

0.43 4.03 

2.22 

2.47 


TABLE B 

Mean effect of potassium and phosphorus 
deficiency and of sodium-calcium 
balance on fructosan 
concentration in 
barley 



Potassium 


Phosphorus 

Sodium/calcium balance 


Leaves 

Stems 


Leaves 

Stems 

Leaves 

Stems 

High 

0.93 

2.02 

High 

0.28 

0.86 

High Na 0.45 

1.01 

Medium 

Low 

0.53 

0.18 

1.19 

0.24 

Low 

1.82 

1.44 

High Ca 0.65 

1.29 















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- 8 


Gregory (12), in an extensive study of mineral 
nutrition of plants found that nitrogen deficiency had no con¬ 
sistent effect on reducing sugars but that the total carbohy¬ 
drate was greatly increased. The effect of phosphorus 
deficiency resulted in higher levels of free reducing sugars 
but total sugar was less affected; while potassium deficiency 
lowers both free reducing sugar and total sugar. 

Richards (16) investigating a similar problem, found 
evidence* supporting Gregory’s observations and confirmed the 
fact that the various mineral treatments affected the ratio 
of sucrose to reducing sugars. Finally, Russell (17) found 
ratios of fructosan to other sugars greatly influenced by the 
various conditions of mineral deficiency, and that in the 
final analysis the fructosan accumulation was controlled 
almost wholly by the concentration of other sugars. 

On the basis of experiments on mineral deficiencies 
it becomes clear that fructosans accumulate under all condi¬ 
tions promoting storage of sugars. 

The studies on carbohydrate content of plants grow¬ 
ing under field conditions is confined almost entirely to the 
observations Barnell (9) (10) made on wheat. Two wheat 
varieties were selected and planted in the fall, and harvested 

* No consistent differences ascribable to potassium were 
found. 



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the following summer. The percentage of sucrose in the plants 
was found to some extent to be a function of temperature, the 
sucrose value increasing with low temperature and falling 
with moderate temperature. During one cold period sucrose 
rose from a value of ,25$ of the fresh weight to a peak, 17 
days later, of 1,55%. The low temperatures were associated 
with more than average sun so that, under frost conditions, the 
rise in sucrose concentration may be due, to some extent at 
least, to the rate of production of sucrose by photosynthesis 
exceeding the rate of utilization in the growth and respira¬ 
tion processes. Temperature conditions resulted in greater 
fluctuations of sucrose content than did hours of sunshine 
when results were analyzed statistically. Barnell (10) con¬ 
sidered sucrose to be the dominant sugar and, except in the 
later stages of growth when the highest value for total sugar 
was observed, glucose exceeded fructose. The values obtained 
by him are as follows: 

Maximum Values of Fructose, Glucose, and Sucrose (% of fresh 

weigFtTT3' j-- --- 

Time of head emergence maximum Glucose - .50% 

Two weeks later " Sucrose - 5 to 4% 

At same time or a 

little later " Fructose exceeds glucose 

until harvest. 

No significant differences were found in sugar con¬ 
centrations of the two wheat varieties except that in the 
later variety the maximum values were reached later. Barnell 




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10 


finally suggests that the sugar stored is not essential to 
filling of the grain, and thus supports the conclusions ad¬ 
vanced by Archbold and others* 

No reference to daily variations of fructosans is 
reported in the literature reviewed* While the work of Curtis 
is included in the references of this report it must be under¬ 
stood that no attempt to estimate the fructosans was made by 
him. Curtis (11) observed that the sugar content of alfalfa 
is greatly influenced by the time of day at which it is cut* 

The observed results indicate differences between afternoon 
and morning cuttings are of the order of 83$ or, in some 
cases, more* These differences were estimated to fall almost 
entirely on the soluble fractions of the plant materials and 
would therefore include all the sugars. Curtis further 
suggests, on the basis of observations made on carbohydrate 
losses during "curing”, that losses of soluble sugars may 
reach proportions of 50$ of the amount known to exist in the 
green material. Woodward, Tisdale, and Shepherd (EO) disagree 
with Curtis, stating that such high losses cannot be real, 
since losses due to respiration could not account for such 
large amounts of sugar* It has already been pointed out, 
nevertheless, in references to the work of Archbold and 
others, that losses of sugar from plant tissues have been 
observed to be large* 

It would seem from the foregoing that there is a need 
for more evidence from experiments with plants to determine 


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11 - 


more exactly the magnitude of these changes as related to 
time factors, soil, and weather effects, and kind of crop 
concerned. 

At present, only the all-over quantitative changes 
in sugar concentrations appear to be known. Archbold and 
Datta (7), in their more recent work, have concluded that, 
before much more can be known of the distribution of the car¬ 
bohydrate supply for either growth processes or storage, 
nitrogen metabolism must be considered in relation to carbo¬ 
hydrate synthesis and utilization. This view is further 
borne out by work of Richards (16) who demonstrated that a 
remarkably close relationship exists between respiration rate 
and protein content. 

It may be of interest to note that Archbold is now 
engaged in experiments on the part played by nitrogen in 
determining the distribution of the available carbohydrate 
supply. 


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- 12 


MATERIALS AND METHODS 


Materials 


The barley used in these experiments was grown from 
foundation stock seed of Newal at the University of Alberta. 
Replicate seedings were made on May 3, May 17, May 25, June 3, 
and June 22* 

Each of the first seedings consisted of one 40-foot 
row. This material was used in the preliminary testing of 
methods of analysis. The final planting consisted of 24 
rows, 50 feet long and spaced nine inches apart. Plants from 
this seeding were used for most of the analytical work reported 
in this paper. 

The barley was planted on a field of black loam 
which had been summerfallowed the previous year* The field is 
typical of the agricultural soils in the Edmonton area and 
considered to be very fertile. 

The period from June 22 to September 30 was excep¬ 
tionally wet and very vigorous growth of the barley occurred. 
Lodging occurred at the end of July after considerable wet 
weather. The barley was fully headed 57 days after planting, 
at which time it was approximately 42 inches high. The last 
collection was made 100 days after planting. The late sowing 
and abnormally high rainfall contributed to late maturity, so 








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-13 - 


that even at the end of 100 days many tillers had not 
ripened, although the growth period for Newal grown at Edmon¬ 
ton, on the basis of a seven-year average, is 88 to 90 days. 

Method of Collection 


At weekly intervals, beginning on the eighth day 
after planting and ending after 100 days of growth, single 
plants selected at random over the whole plot were cut with 
shears at ground level until 25 to 150 plants were obtained. 

To measure the effect of time of collection, one collection 
was made at 4:30 in the afternoon and another at 8:30 the fol¬ 
lowing morning. 

The whole plants were immediately brought to the 
laboratory where they were cut into half-inch lengths and 
thoroughly mixed. A forty-gram sample was weighed out and 
placed in 200 cc. of boiling 80$ alcohol. Ten to twenty grams 
for moisture content determinations were placed in a drying 
oven kept at 75 to 80°C. The remainder was placed in large, 
shallow pans and allowed to dry at room temperature. This 
material, when thoroughly dry, was ground in a Wylie mill and 
placed in jars until required. 

Beginning on August 18, weekly analyses for various 
carbohydrates were made on the component parts of the plants 
collected at 4:30 p.m. Heads were separated at the junction 
with the peduncle, leaves detached at the junction with sheaths 



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- 14 - 


and sheaths peeled from the stems. After separation—which 
was done as quickly as possible—the various parts of the 
plants were cut up. Mixed portions of 5 to 10 grams were 
weighed out and placed in boiling 80$ alcohol. 

Notes on height of plants and stage of development, 
together with notes on weather conditions at the time of 
collection, were taken throughout the growth period. Detailed 
records on weather for the entire period appear elsewhere in 
this report. 


Preparation of Material 


As a rule a 40-gram sample of the freshly collected 
and cut material was placed in 200 cc. of boiling 80$ alcohol 
to prevent enzymatic hydrolysis of fructosans and sucrose. 
Since fructosans are not extracted by alcohol, a water 
extraction was carried out on the material which was first 
freed from alcohol. The alcohol was poured off and the plant 
material washed free of alcohol by shaking with several addi¬ 
tions of water. The residual material was transferred to a 
Waring Blendor, sufficient water added to cover the material, 
and cutting in the blendor was carried out for approximately 
five minutes. It was observed that the material was reduced 
to a very finely divided condition. After transfer to a 
wide mouth Erlenmeyer, the cut material was shaken in a 



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15 - 


mechanical shaker for approximately two hours * The result¬ 
ing water extract was transferred to a graduated flask to 
which the residue of the alcoholic extract and washings were 
added after the alcohol had been removed by distillation under 
reduced pressure. The residual material was then washed by 
shaking with small additions of water and decantation of the 
washings into the flask until the required volume was nearly 
attained. The liquid was finally made up to volume and pre¬ 
served by the addition of a little toluene. The extracts 
obtained from the young material were always deeply colored 
by the pigments extracted during water and alcohol treatments. 
Older plants, especially near the end of the growth season, 
did not give such highly colored extracts. 

When analyses were mad© on the dried materials the 
same procedures were used, except for the weight of material 
and final volume of extract. 

Residues after alcoholic and water extractions were 
re-extracted and in no case was there an appreciable amount of 
total sugar remaining in the residue, and the amount which did 
remain was of the same order for different samples. 

Results of some of such re-extractions are shown in 


Table C 




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- 16 - 


TABLE C 

Total sugar content in water extracts of barley 


Age (days) 

1st extraction 

2nd extraction 

Green plants 

($ of green weight) 


8 

1.22 

.03 

29 

1.46 

.03 

54 

2.83 

.04 

Dried plants 

(% of dry weight) 


21 

3.76 

.16 

35 

2.12 

.15 

49 

8.38 

.15 

63 

11.29 

.15 

77 

6.32 

.09 

91 

1.31 

.13 


* Removal of alcohol from the alcohol extract and 
washings presented many difficulties when lack of pressure in 
water taps made the use of water aspirators unreliable. This 
was finally satisfactorily overcome by using a vacuum pump on 
a system consisting of two HgO condensers, a Claissen flask, 
and a large CaClg trap. Continual use of the pump over 
periods of a month or more revealed little dilution of oil in 
the pump. When the vapor of the alcohol and water solution 
was pumped off at 2 to 3 cm. of mercury, reduction in volume 
of some 200 to 300 cc. was possible in three hours or less. 

A fine capillary tub© was used to admit small bubbles of air 
into the solution during progress of distillation in order to 
prevent frothing and bumping. 







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- 17 - 


The moisture and dry weight determinations were dif¬ 
ficult to make accurately because of the rainy conditions 
that prevailed during periods of collection. Plants col¬ 
lected in the morning were, with few exceptions, either 
covered with heavy dew or else with rain which had fallen 
during the night. No attempt was made to dry off excess 
moisture before sampling, since it was essential that analysis 
be made on the plants as soon after collection as possible. 

Any large drops of surplus moisture on the plants were merely 
shaken off. 

Considerable error may arise from this source and, 
in any future work, some attempt should be made to overcome 
this difficulty. 

Methods of Analysis 


A. Reducing Sugars 

The modified Harding and Downs copper reduction 
method was used in all estimations of reducing sugars. 

Separate estimates of fructose were made after oxidation of 
the glucose by hypoiodate. Both procedures are described 
fully by Van der Plank (19). 

Values for reducing sugars in terms of standard .01 
Normal sodium thiosulphate were obtained by using various con¬ 
centrations of chemically pure fructose and glucose, and 






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- 18 - 


plotting the results on a graph. These values were used to 
estimate the reducing power of the plant extracts. Consider¬ 
able testing of the procedure for estimation of sugar was done 
and the results of such testing appear in Tables D and E. 


TABLE D 

Copper-reducing power of glucose and fructose in 
terms of .01 normal sodium thiosulphate 


Sugar 
(mg.) 

Glucose 
(boiling time - 

15 min.) 

Fructose in presence of 
.221$ KJ 

(boiling time - 10 min.) 

van der Plank < 

Observed 

van der Plank 

Observed 

.125 

.52 

mmmm 



..25 

1.06 

1.21 

.81 

.80 

.50 

2.20 

2.30 

1.89 

1.90 

.75 

— 

3.35 

<3X0 <m» 

3.00 

1.00 

4.55 

4.60 

4.08 

4.10 

1.25 

mm me 

5.75 


5.25 

1.50 

6.88 

6.90 

6.41 

6.35 

1.75 

... 

8.10 

... 

7.50 

2.00 

—- 

9.25* 

— 

8.60 

2.25 


10.40* 

mmm* 

-• 

2.50 

— 

11.60* 



2.75 


12.75* 

—- 

— 

3.00 


13.90* 

— — 



* Double volumes of all reagents used. 







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- 19 


TABLE E 

Recovery of glucose and fructose added to 
plant extracts (results expressed as <?<>) 


Recovery 

Extract + sugar Extract Observed Theoretical 

Glucose 


1 

1.50 

1.14 

.36 

.40 

2 

4.18 

2.40 

1.78 

1.88 

3 

3.87 

2.98 

.89 

.80 

4 

1.82 

.85 

.97 

.98 

Fructose 

1 

.76 

.36 

.40 

.40 

•2 

1.58 

1.20 

.38 

.40 


Fermentations using the methods described by Yemm 
(21) were found to be entirely satisfactory and, while cor¬ 
rections for the magnitude of the reducing power due to 
non-fermentable substances were not made on all extracts, 
enough were actually carried out to make this correction pos¬ 
sible* Results of fermentation analysis are shown in Table F. 








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20 - 


TABLE F 

Estimates of non-fermentable substances 
in plant extracts 


Plant age 
(days) 

On green materials 
[io green weight) 

On dried materials 
($ dry weight) 

Charcoal 

cleared 

Uncleared 

Charcoal 

cleared 

Uncleared 

20 

.02 

.04 

.13 

.15 

30 

o01 

.04 

- 

- 

40 

.06 

.04 

.03 

.20 

50 

.05 

.06 

.01 

.24 

60 

o 

- 

.01 

.42 

70 

- 

mm 

- 

— 

80 

— 


.02 

.24 


Methods for clarification of non-hydrolyzed extracts 
involved the use of charcoal in the manner suggested by 
Archbold (1)* Results obtained appear to be in agreement 
with those obtained by Archbold. A few of the observed 
results are included in Table G. 







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21 - 


TABLE G 

Estimates of glucose in water extracts of 
barley before and after clarifi¬ 
cation with charcoal 


Glucose corrected 
for non-fermentable 


Before fermentation After fermentation substances 


Sample 

Uncleared Cleared 

Uncleared 

Cleared 

Uncleared 

Cleared 

1 

1*33 

1.21 

.14 

.04 

1.19 

1.17 

2 

1.07 

1.02 

.16 

.09 

.91 

.93 

3 

1.48 

1.35 

.23 

♦ 07 

1.25 

1.28 

4 

1.01 

.89 

.19 

.03 

.82 

.86 

Amounts 

of charcoal 

used were from .4 

to .6 g* 

per 100 ml. 

of 

solution containing 

4 g. Of 

fresh material per 

100 ml. 



It should be observed that clarification results in lowered 
reducing power, owing to non-fermentable substances but, at 
the same time, results in many instances of increase in value 
for corrected reducing sugars* It has been suggested by 
Archbold (2) that reducing substances other than sugars 
diminish the reducing power of sugars present* 


The methods which were used in routine estimations 
of sugars and which have been referred to, appear to be 
entirely trustworthy if all the conditions are carefully fol¬ 
lowed* It has been observed, however, that the following 
precautions should be emphasized: 

1* All reagent solutions used must be accurately 
and carefully measured. 

2* The heating period of 15 minutes must be ad¬ 
hered to under conditions that remain constant from run to run. 

3. The "holding period" at 1°C. for two hours in 








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one of the steps of hypoiodate oxidation of glucose is 
exceedingly difficult to maintain unless there is available a 
dependable refrigerator. This troublesome factor was success¬ 
fully overcome by placing the flasks in ice and water mixtures 
before placing in the refrigerator. 

4. The capacity of the "sugar reagent" to measure 
reducing power is limited. It was observed that, if the con¬ 
centration of the sugar in the aliquot was such that reduction 
nearly equalled the blank, results were invariably unreliable. 
A difference of at least 1.5 ce. of .01 normal sodium thio¬ 
sulphate should obtain between reduced reagent and the blank. 

5. Addition of charcoal to extracts must be care¬ 
fully carried out. Excesses of charcoal were found to remove 
sugars in rather significant amounts. The amount used should 
just clear the extracts and no more. The amounts used, there¬ 
fore, varied with the amount of pigment present in the solu¬ 
tions and less was used with older plants than with younger 
ones. 

6. Neutralization of acidified extracts is very 
unsatisfactory if end point of neutralization must be made by 
relying on indicator dyes. The difficulty of observing 
indicator color change in opaque solutions is great. Some 
buffer effect was also observed in the hydrolized extracts. 

The use of a portable glass electrode potentiometer is essen¬ 
tial for consistent results. 


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Technique used in the Estimation of Sugars 


1. Free reducing sugars were estimated on clarified 
extracts, using the modified Harding and Downs microcopper 
reagent. Total sugars were estimated after hydrolysis with 
N/5 H 2 SO 4 , no clarification being necessary. 

2. Separate estimations of fructose were made on clari¬ 
fied, non-hydrolyzed and non-clarified hydrolyzed extracts. 

5. After corrections in free reducing and total sugars 

for non-fermentable substances had been made: 

(a) Free reducing sugar minus free fructose s free 

glucose. 

(b) Total reducing sugar minus total fructose s total 

glucose. 

4. The approximate estimate of fructosan was made from 
the excess of fructose over glucose produced by N/5 acid, plus 
a correction of 6 $ glucose occurring in the known fructosan. 

5. Estimates of sucrose as the difference between 
increase on hydrolysis and the fructosan estimate were subject 
to large errors and therefore not reliable (4). 

6 . A second means of estimating sucrose is suggested 
whereby twice the increase in glucose produced on hydrolysis 
equals sucrose. These values will always be too high owing to 
the inclusion of glucose from sources other than sucrose. At 



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24 - 


best they can only be regarded as approximations (4). 

EXPERIMENTAL RESULTS 


As a matter of convenience the experimental results 
obtained in this investigation are divided into four sections: 

Section A - The drifts in sugar and dry matter con¬ 
tent in growing barley from the eighth day after seeding 
to the end of the growing season. 

Section B - The differences in sugar and dry matter 
content in growing barley as related to the time of day 
collections were made. 

Section C - The sugar and dry matter content in the 
stems, sheaths, leaves, and heads of growing barley from 
the 57th to the 85th day after sowing. 

Section D - The losses of sugars from barley plants 
air-dried at room temperatures. 

Seasonal Drift in Sugar and Dry Matter Content 


Section A 

The results for this section appear in Tables I, II, 
and III; and Figures 1 to 5. 

The plentiful supply of moisture during the season 













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25 


resulted in very vigorous growth and before heading many of 
the plants had already lodged* The leaves and stems were 
larger than those normally produced and long before heading 
many of the lower leaves showed yellowing, largely the result 
of excessive shading owing to lodging* On only four of the 
28 collection days was there no rain in the 24 hours preceding, 
and on many occasions it was raining when samples were taken* 
The actual moisture content of the material was difficult to 
obtain* In this section an attempt is made to overcome this 
difficulty, in part at least, by reporting the results as 
the average of the afternoon and morning values and omitting 
the percentages based on green weight in all except a few 
instances* 

Heading was practically complete on the 57th day 
after sowing* From the 29th day to the 57th day 55 inches of 
the 48 inches of the season’s growth took place* Up until 
heading the drifts in the concentrations of all sugars were 
subject to considerable variation* Whereas there was a slow 
increase in total sugar, the free reducing sugars showed a 
gradual increase, reaching their maximum content for the 
season on the 57th day* Despite the great increase in height 
of plants there was no great increase in the percent dry 
matter. Many large tillers were produced* 

The amounts of free glucose exceeded free fructose 
throughout the growth season. 


Concentration increases in one sugar usually were 


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26 


associated with increases in another, so that there was evi¬ 
dent a gradual upward trend in all of them. This was very 
noticeable shortly before heading when increases in all sugars 
were large. Throughout most of the period total fructose 
exceeded total glucose, the maximum concentrations reached 
coinciding with those for the free reducing sugars. 

The fructose produced by acid hydrolysis, after cor¬ 
rections for the amounts arising from sucrose, appears as 
fructosan. Except for the rather low values of this sugar 
obtained on the 36th and 43rd day there was a gradual upward 
trend until the 57th day, when the maximum concentration was 
reached. The low values were difficult to explain. The 
period was marked by large variations in hours of sunshine, 
temperatures, and amounts of rainfall. It rained almost con¬ 
tinuously during July 29, 30, and 31, and there was less than 
one hour of sunshine recorded. The maximum temperature 
readings were almost 10°F. less than the July average. The 
period following was bright and warm. 

The fructosans, at their maximum concentration, 
formed approximately 35$ of the total sugar, but the absolute 
amounts were not large. 

Sucrose was the dominant sugar throughout the whole 
period, reaching its maximum concentration a week or so after 
the fructosan maximum. 

No starch was observed in any of the tissues of the 


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27 


plants until head formation, when positive iodine tests were 
obtained in the head materials. 

In the week following heading, the maximum concen¬ 
trations of all sugars had been reached and, from this date to 
the end of the season, a sharp decrease in the amounts of all 
sugars was clearly evident. The presence of fructosans in 
the plants after the 78th day was doubtful. The drifts in 
the concentrations of all sugars almost paralleled one another, 
until at the end of the growth period only small amounts of 
sugar remained. 

For comparative purposes the following observations 
may be related to periods of growth. 

A. From Sowing to Heading 

1. The total sugar increased slowly in the early 
part of the growth cycle. At the same time there was a 
more rapid increase in dry matter content. 

2. There was a gradual increase in the amounts of 
both free and combined fructose and in glucose. 

3. The sucrose level remained fairly constant. 

4. All sugars, with the exception of the total 
sugar and sucrose, reached their maximum concentration 
at time of heading. 

5. This period marked the time of most active 
growth and, by heading, most structural parts of the 
plants were formed. 

6 . Some of the lower leaves were dead by the time 


heading was complete 



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B» First week after Heading 

1. The maximum concentration of all sugars was 
reached. 

2. Head formation began and was accompanied by a 
rapid increase in dry matter content. 

C. One week after Heading to End of Season 

1. The leaves died and ceased to function as centres 
for assimilation. 

2. The dry matter content increased rapidly. 

3. The concentrations of all sugars decreased 
rapidly. 

4. Most of the starch in the grain was formed 
during this period. 

On the basis of recent conclusions reached by 
workers on this problem, the variable trends of all sugars 
in the early stages of growth are due to the great demands of 
the actively growing plant for primary assimilates, thus 
leaving little for purposes of storage. The leaf surfaces 
which are the most important sources for photosynthate in the 
early stages were not large enough to do more than supply 
immediate demands of growth. It is only when the plants 
become fully developed that demands are satisfied and an 
excess is produced for condensation to sucrose and fructosan. 
This condition should normally exist when growth of the stem 





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- 29 - 


is nearly complete. This is borne out in the observed con¬ 
centrations of stored sugars found at or near heading. If 
the view that stored sugar represents a simple balance of 
demand over supply is accepted, then active and vigorous 
growth should result in relatively lower concentrations of 
stored sugars. The results of the present experiments appear 
to be in agreement with these conclusions, since they do not 
give the high values observed in plants growing under condi¬ 
tions known to favor sugar storage. 


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30 


TABLE I 
Field notes 


Date of 
collection 

Stage of growth 

Dry 

matter 

M) 

Weather at collection time 

June 

30 

1 leaf 

10.9 

Cloudy - trace rain 

July 

1 

1 leaf 

10.5 

Cloudy - rain overnight 

July 

7 

2 leaves 

11.1 

Cloudy - trace rain 

8 

2 leaves 

10.7 

Cloudy - raining 


14 

3 to 5 leaves 

10.2 

Fair - warm 


15 

3 to 5 leaves 

11.0 

Shower - warm 


21 

Flag leaf 

14.9 

Bright - no rain 


22 

Flag leaf 

13.8 

Bright - no rain 


28 

(Some lodging - 

17.8 

Showery - trace rain 


29 

( awns showing 

14.0 

Cloudy - rain overnight 

Aug. 

4 

Head emergence 

18.4 

Bright - no rain 


5 

Head emergence 

15.3 

Bright - rain overnight 


11 

(Almost headed 

21.6 

Very cloudy - shower 


12 

(Almost headed 
( badly lodged 

19.8 

Bright - rain overnight 


18 

Lower leaves) 

25.6 

Cloudy - cool - damp 


19 

yellowing ) 

23.1 

Heavily overcast - no rain 


25 

(Dough stage;end 

28.8 

Cloudy - shower in p.m. 


26 

( of max. growth 

26.7 

Cloudy - heavy dew 

Sept. 

1 

Many lower ) 

30.3 

Heavy clouds - no rain 


2 

leaves dead) 

28.2 

Cool - heavy dew 


8 

(All lower 

34.4 

Cloudy - cool; trace rain 


9 

( leaves dead 

35.1 

Cool - bright 


15 

Heads ripening 

46.3 

Cloudy - cold - damp 


16 

Heads ripening 

43.1 

Raining - cold 


22 

(Plants nearly 

47.4 

Cloudy - cool - no rain 


23 

( ripe; some 
( tillers green 

47.0 

Cloudy - cool - dew 


29 

Harvest 

52.9 

Cold - rain - snow 


30 

Harvest 

52.3 

Gold - rain and snow 





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31 


TABLE II 

Composition of aerial parts of barley during 
growth (sugar concentrations expressed 
as percent green weight) 


Collection date 

Total sugar 

Reducing sugar 

Fructosan 

June 30 

1*08 

.31 


July 7 

.85 

.36 

-- 

14 

1*03 

.28 

.01 

21 

1*31 

.52 

.35 

28 

1*21 

.63 

.21 

Aug* 4 

1.34 

.68 

.14 

11 

1.52 

.93 

.63 

18 

3.44 

1.16 

1.20 

25 

5.22 

1*28 

1.09 

Sept. 1 

5*07 

1.45 

— 

8 

3.85 

1.16 

.08 

15 

2.05 

.55 

oa 

22 

1.65 

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29 

1.06 

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Composition of the aerial parts of barley during growth 
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32 



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Figure 1 

The drifts in sugar and dry matter 
content in growing barley 

Note: Weather data shown are those which prevailed in 
the day preceding collection. 

Dry matter as percent of the green weight. 























34 



Figures 2 and 5 

Composition of barley during growth. 

Sugar concentrations as averages of after¬ 
noon and morning collections 










SUGARS % OF DRY WT. 


35 



Figure 4 

Composition of barley during growth. 


Free glucose and free fructose as 
averages of afternoon and 
morning collections 









SUGARS % OF DRY WT. 


36 



Figure 5 

Sugar eonoentrations in growing barley 
as averages of morning and after¬ 
noon collections 


Glucose and fructose as increases 
after acid hydrolysis 














' 








. 



















- 37 


Section B 


The investigation discussed in this section was made 
in an attempt to estimate the extent of sugar and dry matter 
changes following periods of darkness. At present there is 
considerable evidence (8) that losses of stored sugars are 
not due to the demands of growth but are apparently related 
to respiration*. 

Some striking results were obtained in the prelimi¬ 
nary work when sugar concentrations were estimated in plants 
growing under dull and wet conditions. Collections were made 
at approximately 1:30 p.m. on three different dates and the 
data presented in Table IT. 


TABLE TV 

Sugar concentrations expressed as percent 
of green weight 


Collection 

date 

Age 

(dy.) 

Total 

sugar 

Reducing 

sugar 

Sucrose 

Total 

glucose 

Total 

fructose 

June 9 

15 

1.22 

.39 

.10 

.44 

.78 

June 12 

18 

.44 

.15 

.03 

.14 

.30 

June 16 

22 

.26 

.06 

— 

.21 

.05 


* Losses have been attributed to respiration activities 

although at present no conclusive evidence has been obtained 
as to the actual fate of stored sugars (6). 








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- 38 - 


The period from June 4th to 9th was exceptionally 
warm and the hours of sunshine recorded for each day were 
much higher than the average value for the month* In the 
following week, 4.17 inches of rain fell and there was no 
direct sunshine from June 13 to 16. The very sharp and well 
defined decreases in concentrations of all sugars were clearly 
related to the prevailing conditions of weather, and it would 
appear that the low hours of sunlight and low temperatures 
influence sugar concentrations because losses due to respira¬ 
tion are not replaced by photosynthetic activity. 

The remainder of the discussion in this section is 
concerned with the barley planted on June 22, the data appear¬ 
ing in Table J and Figures 6, 7, and 8. 

The dry matter content in the morning collection 
was, with few exceptions, lower than that of comparable after¬ 
noon collections. The largest differences were noted when 
the plants were young and growing rapidly, but for reasons 
already considered, the difficulties involved in making mois¬ 
ture determinations made actual estimates of dry matter 
questionable. A definite relationship between dry matter dif¬ 
ferences and age was observed to exist. After time of heading, 
the morning and afternoon differences in dry matter content 
decreased with increase in age. 

Differences in sugar concentration were not clearly 
evident. There is some reason to suggest that the total 
sugar values were higher in the afternoon, but the fact that 


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39 - 


TABLE V 

Sugar content of barley expressed as percent 
of dry weight, as related to time of 
day collections were made 

A = afternoon collection 
B = collection following 
morning 


Age 

of 

plant 

Total 

sugar 

Sucrose 

Fructose 

after 

hydrolysis* 

Free 

glucose 

Free 

fructose 

A 

B 

A 

B 

A 

B 

A 

B 

A 

B 

8 

13.76 

6.29 

11.56 

4.20 

4 0 95 

1.52 

1.65 

1.81 

1.38 

.86 

15 

7.66 






1.80 




22 

10.78 

8.64 

7.24 

6.56 

4.12 

3.18 

1.67 

1.36 

1.37 

.82 

29 

11.07 

6.96 

2.68 

4.34 

5.84 

1.67 

2.95 

2.54 

.94 

.58 

36 

8.09 

7.00 

3.02 

2.00 

2.87 

1.79 

2.70 

3.14 

1.01 

1.07 

43 

8.48 

7.32 

4.14 

2.10 

2.28 

2.35 

2.39 

3.00 

1.74 

.92 

50 

7.31 

7.37 

1.28 

— 

3.01 

2.47 

1.58 

3.58 

2.08 

1.77 

57 

14.10 

14.16 

5.54 

4.52 

6.41 

7.35 

3.32 

2.90 

1.60 

1.65 

64 

17.78 

19.92 

9.32 

12.30 

8.57 

9.13 

2.88 

3.10 

1.67 

1.54 

71 

20.26 

14.15 

19.66 

12.42 

4.95 

3.54 

3.17 

2.34 

2.31 

2.06 

78 

11.80 

10.34 

6.88 

8.32 

4.91 

3.05 

1.86 

1.59 

1.69 

1.54 

85 

4.58 

4.59 

4.96 

4.12 

1.07 

1.13 

.55 

.89 

.49 

.52 

92 

3.04 

3.94 

1.68 

— 

— 


.17 

-- 

— 

— 

99 

2.31 

1.70 

1.76 

1.52 

—— 

—— 

.13 

.02 

•*— 

—— 


* Total increase in fructose after acid hydrolysis. 


some of the afternoon values were lower than those for the 
morning makes acceptance of this open to question. The only 
real differences were found in the period of one and one-half 
months immediately after seeding. Near harvest the differences 
were much smaller and were not consistent. 

The free glucose estimations varied so greatly that, 
with the exception of the first few observations, no consistent 
differences were noted. The free fructose concentrations, on 













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40 


the other hand, showed real differences and, with only one or 
two exceptions, the afternoon values exceeded those for the 
morning. The differences in the early stages of growth were 
much greater than those noted in the periods approaching 
maturity. Increases in fructose after acid hydrolysis were 
higher in plants collected in the afternoon, and followed 
drifts much like those for free fructose. The differences 
again appeared to he significant. The fruetosan values are 
directly related to fructose increases following hydrolysis 
so that they too follow much the same concentration changes in 
relation to the time of day at which collections were made. 

The sucrose estimates are at best only approximate 
and are always too high, since glucose from sources at present 
undefined is included (4). The data on sucrose showed no real 
differences, although amounts vary within wide limits. 

Great, variation in weather conditions occurred 
throughout the growth period. From evidence obtained on the 
effect of hours of sunshine and temperature on level of sugar 
concentrations in plant material investigated in the early 
part of this work, it would appear that rather large variations 
must accompany weather changes. If, for example, cloudy con* 
ditions prevailed previous to the 4:30 p.m. collection and 
clear bright weather followed until nightfall, no normal esti¬ 
mate for that day could be expected. The observation for the 
morning collection on the other hand would be dependent upon 
the conditions which obtained from dawn to 8:30 a.m. If 


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41 


storage and formation of sugars is greatly influenced by the 
hours of sunshine during the period before nightfall and the 
period preceding 8:30 a.m., the effect of darkness becomes 
obscured. This difficulty could be overcome in future work 
by taking the "afternoon" collections at sundown and the morn¬ 
ing ones somewhat earlier than 8:30 a.m. As the data appear in 
this report, little can be known of the effect of the hours of 
darkness, since some hours of daylight occurred in the inter¬ 
val between afternoon and morning collections. The differences 
between morning and afternoon collections, nevertheless, did 
indicate a large daily variation in sugar content. 


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42 - 



Figure 6 

Dry matter and total sugar content in 
growing barley as related to time 
of day collections were made 















43 - 


K- 



Figure 7 

Free glucose and free fructose content of 
growing barley as related to time of 
day collections were made 










44 



Figure 8 

Sucrose and total fructose in growing 
barley as related to time of day 
collections were made 










45 - 


Section C 

This portion of the investigation fell considerably 
short of original expectations. The analysis of the separate 
parts of the whole plant was undertaken so that an estimation 
of the level and the time of occurrence of maximum concentra¬ 
tions of sugars could be made* The results obtained indicate 
that the maxima had either been reached or nearly so in all 
parts at the time the first analysis was made and, as a con¬ 
sequence, only the "falling phase" of sugar trends was ob¬ 
served. The data are presented in Tables 71 and VII* 

In all parts of the plant there was in increase in 
the dry matter content, the larger increases occurring in the 
stems, leaves, and heads. As the first observations were made 
at approximately the time of full heading, a good picture of 
the dry matter increase in the head was obtained. Dry matter 
changes in sheaths were small. Some doubt may be cast on the 
results obtained for the leaves, as many of the lower ones, 
especially in the later periods of growth, had either dropped 
in the field or were lost in handling. At best the results 
should be considered only as approximate estimates. 

The large increases in the dry matter content of 
whole plants is closely related to increases of dry matter in 
the heads. By the time heading was complete it may be assumed 
that most of the structural parts of the plant had been formed 
so that changes of dry matter in all parts but the heads were 
caused by loss of water and tissue degradation, associated with 



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46 


TABLE VI 

Sugar content of steins, sheaths, leaves, 
and heads of barley, expressed 
as percent of dry weight of plant 


Date of 
collection 

age 

Stems 

Sheaths 

Leaves 

Heads 

Total 

Whole 

plant* 

Dry matter 

as % of 

fresh weight of whole plant 


Aug. 18 

57 

<■»«» 


•mwm* 




25 

64 

39.3 

13.6 

18.8 

38.4 


28.8 

Sept. 1 

71 

37.9 

13.1 

16.1 

33.4 


30.3 

8 

78 

35.5 

13.6 

13.2 

37.7 


34.4 

15 

85 

35.3 

13.6 

7.2 

45.0 


46.3 


Total 

sugar as % dry weight of plant 



Aug. 25 

64 

9.01 

1.86 

2.30 

4.93 

18.10 

17.78 

Sept. 1 

71 

8.61 

1.80 

1.96 

4.51 

16.88 

20.26** 

8 

78 

5.21 

2.01 

1.55 

2.08 

10.85 

11.80 

15 

85 

1.74 

.64 

.21 

1.31 

3.90 

4.58 


Free glucose 

as % dry weight of plant 



Aug. 25 

64 

1.86 

.40 

.47 

.33 

3.06 

2.88 

Sept. 1 

71 

2.04 

.40 

.48 

.54 

3.46 

3.17 

8 

78 

1.32 

.52 

.38 

.13 

2.35 

1.86 

15 

85 

.29 

.13 

.06 

.10 

.58 

.55 

Free fructose 

as % dry weight of plant 



Aug. 25 

64 

.86 

.18 

.23 

.22 

1.49 

1.67 

Sept. 1 

71 

1.53 

.38 

.32 

.21 

2.34 

2.31 

8 

78 

1.22 

.39 

.27 

.10 

1.98 

1.69 

15 

85 

* 51 

.15 

.04 

.09 

.79 

.49 

Glucose 

} as net 

increase after hydrolysis 




as 

% dry weight of 

plant 




Aug. 25 

64 

2.17 

.55 

.65 

1.45 

4.82 

4.66 

Sept. 1 

71 

2.45 

.52 

.57 

1.49 

5.03 

9.83 

8 

78 

.75 

.37 

.77 

.47 

2.36 

5.30 

15 

85 

.37 

.06 

.07 

.90 

1.40 

3.03 










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47 - 


TABLE VI (continued) 


Date of Whole 

collection Age Stems Sheaths Leaves Heads Total plant * 


Fructose as net increase after hydrolysis 
as % dry weight of plant 


Aug. 

25 

64 

4.11 

.73 

.96 

2.94 

8.74 

8.57 

Sept, 

, 1 

71 

2.59 . 

.55 

.58 

2.27 

5.99 

4.95 

8 

78 

1.92 

.75 

.13 

1.38 

4.18 

4.91 


15 

85 

.21 

.25 

.04 

.24 

.74 

1.07 


Fructosan as % dry weight of plant 


Aug. 

25 

64 

2.21 

.20 

.35 

1.70 

4.46 

4.45 

Sept. 

1 

71 

.16 

.04 

,01 

.89 

1.10 

— 

8 

78 

1.33 

.44 

— 

1.04 

2.81 

1.79 


15 

85 

.21 

.21 


— 

.42 

— 



Sucrose as 

% dry weight of plant 



Aug. 

25 

64 

4.34 

1.10 

1.30 

2.90 

9.64 

9.32 

Sept, 

1 

71 

4.90 

1.03 

1.14 

2.98 

10.05 

19.66** 

8 

78 

1.51 

.71 

1.53 

.94 

4.69 

6.68 


15 

85 

.74 

.12 

.13 

1.79 

2.78 

4.96 


Reducing sugar as % dry weight of plant 


Aug. 

25 

64 

2.72 

.58 

.70 

.55 

4.55 

4 0 55 

Sept 

. 1 

71 

3.57 

.67 

.81 

.75 

5.80 

5.48 

8 

78 

2.54 

.91 

.65 

.23 

4.33 

3.55 


15 

85 

.81 

.33 

.10 

.18 

1.42 

1.04 


* Obtained on the whole plants on the corresponding dates of 
collection. 

**The apparent over-estimation of the increase in glucose 
after hydrolysis gives: 

1. Too high value for sucrose. 

2. Low value for fructosan. 

3. Too high value for total sugar. 










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48 


TABLE VII 

Sugar and dry matter content in the stems, 
sheaths, leaves, and heads of barley 


Date of Age of 


collection 

plant 

Stems 

Sheaths 

Leaves 

Heads 

Dry matter of the part as % of green 



weight of 

part 



Aug. 18 

57 

32.1 

24.2 

19.2 

30.8 

35 

64 

30.0 

26.6 

24.8 

39.6 

Sept. 1 

71 

25.7 

24.6 

22.9 

39.8 

8 

78 

27.4 

26.4 

28.2 

49.0 

15 

85 

32.6 

29.6 

55.0 

53.8 


Total sugar as % of dry matter of part 


Aug. 

18 

57 

31.04 

17.11 

9.74 

12.08 

25 

64 

22.93 

13.68 

12.26 

17.37 

Sept 

. 1 

71 

22.72 

13.25 

12.18 

13.92 

8 

78 

14.67 

14.81 

11.77 

5.51 


15 

85 

4.94 

5.04 

2.87 

2.92 


Free glucose as % of dry matter of part 


Aug. 18 

57 

10.22 

3.06 

2.60 

3.02 

25 

64 

4.73 

2.94 

2.50 

1.16 

Sept. 1 

71 

5.37 

2.97 

3.01 

1.66 

8 

78 

3.72 

3.86 

2.90 

.34 

15 

.85 

.83 

1.07 

.80 

.22 


Free fructose 

as io of 

dry matter 

of part 


Aug. 18 

57 

2.04 

2.27 


2.34 

25 

64 

2.20 

1.35 

1.21 

.76 

Sept. 1 

71 

4.05 

2.15 

2.01 

.65 

8 

78 

3.43 

2.84 

2.06 

.27 

15 

85 

1.46 

1.22 

.58 

.19 


Glucose as net 

increase after hydrolysis 



as °/o of dry matter of part 



Aug. 18 

57 

3.13 

5.58 


2.11 

25 

64 

5.53 

4.05 

3.47 

5.10 

Sent. 1 

71 

6.46 

3.94 

3.54 

4.60 

8 

78 

2.12 

2.62 

5.82 

1.25 

15 

85 

1.06 

.49 

.91 

1.99 











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49 


TABLE VII (continued) 


Date of Age of 

collection plant_Stems Sheaths Leaves Heads 


Fructose as net increase after hydrolysis 
as jo dry matter of part 


Aug. 18 

57 

15.65 

6.20 

wa> mo 

4.61 

25 

64 

10.47 

5.34 

5.08 

10.35 

Sept. 1 

71 

6.84 

4.19 

3.62 

7.01 

8 

78 

5.40 

5.49 

.99 

3.65 

15 

85 

1.59 

1.96 

.59 

.54 


Fructosan as °/o dry matter of part 


Aug. 18 

57 

14.27 

.71 


2.85 

25 

64 

5.63 

1.47 

1.84 

5.99 

Sept. 1 

71 

.43 

.29 

.09 

2.75 

8 

78 

3.74 

3.27 

-- 

2.74 

15 

85 

.61 

1.68 

-- 

— 


Sucrose 

as io dry matter of part 



Aug. 18 

57 

6.16 

11.16 

OWQMI 

4.22 

25 

64 

11.06 

8.10 

6.94 

10.20 

Sept. 1 

71 

12.92 

7.88 

7.08 

9.20 

8 

78 

4.24 

5.24 

11.64 

2.50 

15 

85 

2.12 

.98 

1.82 

3.98 


Reducing sugar as % dry weight of plant 


Aug. 

18 

57 

12.26 

5.33 

2.60 

5.36 

25 

64 

6.93 

4.29 

3.71 

1.92 

Sept, 

. 1 

71 

9.42 

5.12 

5.02 

2.31 

8 

78 

7.15 

6.70 

4.96 

.61 


15 

85 

2.29 

2.59 

1.38 

.39 









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50 


senescence. 

The most significant results were obtained for the 
dry matter content of the parts as related to that of whole 
plants. The distribution of the dry matter of whole plants is 
briefly summarized in Table VIII. 

TABLE VIII 

Dry matter in the parts as percent of 
dry weight of the whole 


Part 

57th day after seeding 

85th day after seeding 

Stems 

39.3 

35.2 

Sheaths 

13.6 

12.6 

Leaves 

18.8 

7.2 

Heads 

28.4 

45.0 


The changes in dry matter in stems, sheaths, and 
leaves were associated, for the most part, with tissue degra¬ 
dation which occurred during the period of ripening. The 
increase in the dry matter in the head, on the other hand, 
was directly related to the accumulation of starch and other 
materials in the developing grain. At the beginning of the 
study stems formed the larger portion of total dry matter in 
the plant, while near the end heads formed 45$ of the whole. 

In all parts, except heads, two factors responsible for dry 
matter changes were: first, the loss of sugars from the 
tissues; and second, the changes which are related to dehydra¬ 
tion and the transformation that living tissues undergo in 





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51 


the ripening period. Archbold (5) (7) has found no evidence 
of breakdown of any tissue during growth for purposes of sup¬ 
plying the developing grain with starch-building materials, 
so the changes that do occur are associated only with senescence. 

It is clearly evident that the change in dry 
weight is the result of dehydration and death of tissues, at 
least insofar as the stems, sheaths, and leaves are concerned. 
More information would have been obtained had the analysis 
been carried out on a "per plant" basis. The results would 
have shown the actual quantitative changes in dry matter and 
sugar content instead of the percentage changes, which indi¬ 
cate relative rather than absolute quantities in each part. 

The estimations of the various sugars were made for 
each part of the plant and for the whole plant. When a con¬ 
sideration is made of the possible sources of error in sampling, 
and of the errors which result from carrying out so many 
separate steps in the actual analysis, it is concluded that 
reasonably good agreement was obtained in all but a few cases. 

The results for the whole plants collected on September 1st 
appear questionable. 

Most of the sugar was found in the stems and the 
heads, and this was especially true in the earlier stages of 
growth. The concentrations of sugars in the stems and heads 
had reached their maximum levels at or near the time heading 
was observed to be complete. As nearly as can be determined 
the maximum sugar levels for sheaths and leaves had been 


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52 - 


reached earlier. 

Throughout the period the total sugars in the stems 
was about twice that found in the heads, and approximately 
four times the observed values for leaves and sheaths. The 
order of the contribution of sugar each part made to the plant 
was therefore: 

Stems, approximately 50% 

Heads, approximately 25% 

Sheaths and leaves, approximately 25%. 

Individual results varied somewhat but the above is 
a fair approximation. 

In the 28-day period there was a marked decline in 
concentration of the total sugars in both stems and heads, and 
this decline was closely related to the changes in total 
sugars estimated for the whole plants. The trends in leaves 
and sheaths were not so obvious except near the end of the 
period. 

The greater part of the free glucose observed in the 
plant is contributed by the stems. The amounts found in the 
other parts were much lower and, on the whole, there was little 
difference between the concentrations found in various parts. 
The same relationship exists for free fructose, except that 
all parts contain somewhat lesser amounts. The lowest con¬ 
centrations were found in the heads. 

The values for glucose and fructose, as increases 
after hydrolysis, show definite trends, falling steadily from 
the 57th to the 85th day after seeding. The higher concentre- 


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53 - 


tions were again observed to occur in the stems and heads, 
but there were some (differences in the amounts found in the 
leaves and sheaths. 

When reference is made to the sucrose levels in the 
sheaths and leaves it is apparent that a larger increase in 
glucose observed in the leaf material would be expected. The 
fructose differences in leaf and sheath material are less 
obvious, since the larger amounts of fructosans in sheaths 
tend to obscure the effect of the increase of fructose from 
sucrose. 

The fructosan concentrations in all parts of the 
plant varied greatly. There was good evidence that the smal¬ 
lest amounts occurred in the leaves and the greatest in the 
stems, especially at the time active growth of the stem 
ceased. The heads contained significant amounts, and when the 
fructosans in the stem are considered, practically all the 
fructosans in the plant were accounted for. It is difficult 
to say just what the general trend was from the few values 
which were obtained. Generally, the drifts in the whole 
plant are related to the drifts in the parts. It appears 
that the maximum concentration reached occurred at the time 
heading was complete and that, in the period following, the 
amounts decreased rapidly. 

The sucrose concentrations followed trends similar 
to those for other sugars. The only significant fact ob¬ 
served was that the sucrose formed the highest proportion of 



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V .C’C SBV7 Vo vies 


54 - 


the total sugar in almost all parts of the plant and through¬ 
out the period could be considered the dominant sugar. 

Of all the sugars considered, the free reducing 
sugars were subject to the least variation. This fact was 
noted in other parts of this investigation. 

While the period considered is rather short, never¬ 
theless a few general conclusions can be drawn. Up to the 
time of heading and for a short time after, stem constitutes 
the larger portion of the dry matter of the whole plant. In 
the later stages of growth, especially during the ripening 
period, the heads contain almost half the total dry matter. 

In the period following heading, the stems and 
heads contain almost 75$ of the total sugars found in the 
plants and generally show the most marked changes in both dry 
matter and sugar content. When the various sugars in the 
plant are at their maximum the stems contain the highest pro¬ 
portion—as much as 50$ of the sugars. 

The sheaths and leaves contained less sugar than did 
the ears , and the variation in concentration as related to 
the age of the plant did not follow drifts which were as 
obvious. At no time was there much more than 25$ of the total 
sugar ascribable to leaves and sheaths. This, of course, 
would not be true in the early periods of the growth cycle. 

Any changes in sugar concentrations which occurred in either 
of these parts took place in the latter part of the period 
under consideration. 


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55 


All parts of the plant at the end of the Q5th day- 
contained relatively small amounts of sugar. Sucrose was 
the dominant sugar and free fructose was observed to be pre¬ 
sent in the smallest amounts. 

As nearly as can be determined the maximum concen¬ 
tration of sugars was reached in the following time order: 
leaves, sheaths, stems, and heads. This is, of course, just 
what would be expected. Concentration trends of all sugars 
in all parts of the plant were related to the sugar concentra¬ 
tions in the whole plant and appeared to be influenced very 
markedly by the stage of development of the plant, this being 
especially true during the latter stages of the plant’s growth. 

Section D 


The data for this section are presented in Figures 
9 and 10. All sugar values are expressed as percent of the 
weight of the material observed after drying for S4 hours in 
an oven maintained at 55 to 60°C., this low temperature being 
necessary to avoid fructosan destruction. 

A comparison of the seasonal concentration drifts 
of the total sugar estimated in both green and dried material 
indicated close agreement. The total sugar values in dried 
materials were much lower but closely paralleled correspond¬ 
ing values for total sugar in the green materials. The 
maximum total sugar content after drying of the plants was 



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, so i• iiq .o;'s to toil--: iSvit jxxctooo issctotf hi/o/lx^x 


56 - 


11.95$, which was almost 37$ lower than the maximum concentra¬ 
tion before drying. Larger losses occurred with young plants 
which had been growing rapidly. With increasing age of the 
plants losses were not as pronounced. No "average" losses 
could be estimated but an approximation of from 25 to 45$ 
would include most of the individual values determined. 

Concentrations of reducing sugars after drying 
showed that losses varied with age of plant, the greater dif¬ 
ferences occurring when young plants were dried. In these, 
there was a marked reduction in free glucose resulting in the 
presence of larger amounts of free fructose than of free glu¬ 
cose. In the older material the losses of both of these 
sugars appeared to be less evident, so that amounts remaining 
were as high as 80$ of the original values. 

Very large losses of sucrose were noted and, since 
this sugar made up the greater portion of the total sugar in 
the plants before drying, its disappearance during the drying 
period accounted for most of the sugar lost. 

Fructosan losses did not appear to be large, the 
only obvious differences occurring in the younger plants. 

The losses in reducing sugars that actually occurred 
during drying are difficult to determine as sucrose inversion, 
which is broight about by enzymatic activity during drying of 
the plant, would increase the concentration of both glucose 
and fructose (1). The demands of respiration before death of the 



,£ •' ' ’ c • “ * 

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r. . : ;v V ' .• ’v.; .v- o . '.J :v, ,v.v ov r \ O',, t 1 o 

. ( 


57 


plants, however, reduce the amounts of the latter sugars so 
that the amounts remaining would depend upon the extent of 
sucrose inversion and the magnitude of the demands of respira¬ 
tion. The results indicated lowest concentrations of reducing 
sugars to be associated with the smaller amounts of sucrose 
originally present. This phenomenon would also account for the 
relatively higher concentrations of reducing sugars observed 
in plants that contained large amounts of sucrose before dry¬ 
ing. 

It was concluded from the observations on sugar 
losses that occurred during drying of plant material that the 
most labile sugars were glucose and sucrose, whereas fructose 
and fructosans were the least affected. The levels of reduc¬ 
ing sugars in dry material were related to concentrations of 
sucrose originally present. The over-all losses in total 
sugar, due to drying of the plants, was from 25 to 45$. 


■ - t 

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58 - 


TABLE IX 

Composition of dried barley plants 
(Results expressed as fo dry weight) 


Free 

Age of Total reducing Free Free 

plant sugar sugar Sucrose Fructosan glucose fructose 


8 

4.05 

1.22 

15 

2.55 

.93 

22 

3.44 

.83 

29 

4.10 

.93 

36 

2.39 

.48 

43 

1.94 

.52 

50 

5.55 

1.66 

57 

11.03 

3.28 

64 

11.95 

3.80 

71 

10.05 

3.08 

78 

7.17 

2.58 

85 

5.21 

1.64 

92 

2.17 

.94 

99 

.64 

.28 


3.92 

... 

2.76 

— 

2.68 

— 

2.08 

1.24 

1.34 

.57 

.46 

1.09 

1.56 

2.66 

6.12 

1.86 

4.82 

3.80 

6.36 

.70 

4.52 

.07 

2.98 

— 


.34 

.88 

.04 

.89 

— 

.94 

.17 

.76 

— 

.55 

.11 

.41 

1.02 

.64 

1.89 

1.39 

2.22 

1.58 

1.84 

1.24 

1.31 

1.27 

.87 

»77 

.64 

.30 

.15 

.13 








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- 59 - 



Figure 9 

Sugar content in dried barley plants 

Results as averages of afternoon 
and morning collections 











60 



Figure 10 

- Sugar content of dried barley plants 

Free glucose, free fructose, and sucrose 
results as averages of afternoon 
and morning collections 





























61 - 


DISCUSSION AND SUMMARY 


Quantitative determinations of sugars in growing barley, 
with the object of studying seasonal changes and drifts, were 
carried out. Relationships were found among concentrations of 
stored sugars, stage of plant development, and growth rate. 
Weather conditions seemed to be related to rate and extent of 
sugar accumulation. 

When growth rates were high no large accumulations of 
sugars were observed, although during this particular period 
most of the vegetative parts of the plants were formed. Sharp 
increases in all sugar concentrations were noted only when 
active stem elongation was near completion. Maximum concen¬ 
trations of all sugars occurred at or near heading. It would 
seem that in the early periods of the season demands of 
growth processes utilized most of the available assimilate 
and that only after full growth was nearly attained was there 
a surplus of photosynthate produced for purposes of storage. 

Sucrose, always the dominant sugar, rose and fell 
with the other sugars and there appeared to be no evidence 
that the rise in one was associated with the fall in another. 

Any changes in ratios were apparently due to a relatively 
faster increase or decrease of one or the other. Concentra¬ 
tions in all sugars rapidly declined soon after heading until 
at harvest only small amounts remained. 



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62 


The conclusions reached by Archbold (5), Archbold 
and Datta (7), and Archbold and Mukerjee (8) emphasize the 
point that sugar storage is the result of a simple balance of 
supply over demand. The observations and results of this in¬ 
vestigation support this view. 

The seasonal drifts in the concentrations of all 
sugars, as observed in this work, are in general similar to 
those reported, but some differences were apparent. 

The observed amounts of fructosans were lower, while 
the estimations of sucrose were higher, except for Barnell's 
(9) (10) values for wheat. The proportion of fructosans to 
total sugar at the time the maximum concentrations were noted 
was 35$, as compared with reported values as high as 60$ (4). 
Free reducing sugar exceeded fructosan concentrations. The 
presence of fructosans in leaves was not clearly demonstrated. 

The drifts of sugar concentrations in each plant 
component were similar to those observed in whole plants, but 
had different time relations. As nearly as could be determined 
the maximum concentrations of sugars occurred in the following 
time order: leaves, sheaths, stems, and heads. During the 
period considered, the stems were found to contain approximately 
50$ of the sugars found in the plants; heads 25$; and sheaths 
and leaves the remainder. The most apparent decreases in 
sugars during ripening of the plants occurred in heads and 
stems, but near the end of the season large losses from all 
parts were evident. 


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- 63 - 


Losses of sugars from the stems, sheaths, and leaves 
were related to senescent changes in these parts. Fructosan 
concentrations were greatest in the stems and young heads and 
account for most of the fructosan in the plant. Amounts of 
fructosans in all parts of the plant were lower than those 
recently reported by others (4). 

The attempts to obtain quantitative evidence with 
respect to overnight losses in sugars were not very successful, 
largely because conditions were extremely variable* The few 
real differences obtained indicated that free fructose and 
possibly fructosans were affected. The values obtained for 
the other sugars were so inconsistent that no conclusions were 
possible. No reference to diurnal changes in fructosan in 
growing plants was found in the literature, but the results 
of an investigation on sugar concentration variations in 
alfalfa as related to time of day, reported by Curtis (11), 
indicated that large overnight losses were possible. On the 
average, afternoon cuttings were 83 % higher in carbohydrate 
and 19$ higher in dry matter yield than were the morning cut¬ 
tings. 

The dried material contained considerably less sugar 
than did the corresponding samples of green material. The 
observed loss of between 85 and 45$ was found to fall on the 
free glucose and the sucrose fractions. Fructosan losses 
were larger in younger than in older plants, although the 
all-over decreases noted on drying were less than for sucrose 


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64 - 


or free glucose. Decreases in reducing sugars were not evident 
in plants which originally contained high concentrations of 
sucrose. The possibility of sucrose inversion to free reduc¬ 
ing sugars was suggested. 

This preliminary survey of some of the quantitative 
variations in fructosan accumulation in barley indicates that, 
under normal conditions of growth, there is an increase of 
fructosan in the whole plant, at least until stem elongation 
is complete. Subsequently the amount falls as the total sugar 
declines and the grain ripens. It was evident that the accumu¬ 
lation of fructosans did not occur until conditions which pro¬ 
mote sugar storage were present. The ultimate maximum level 
of fructosan was associated with the concentrations of the 
other sugars, while the increase with time was due to increas¬ 
ing sugar content. On the basis of these observations and 
from the conclusions reached by Archbold et al, it became 
clear that fructosans only accumulate under conditions which 
promote sugar storage. Furthermore, sufficient evidence has 
now been obtained to reveal the fact that the accumulation 
and fate of stored sugar, Including fructosans, is not the 
result of simple, easily understood physiological processes 
or the result of any one environmental factor, but rather a 
complex, interrelated combination of all the many factors 
which modify and determine the normal growth of plants. 

Therefore, for the present and in the absence of 
much more detailed and precise information, it can be stated 



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65 - 


that the possibility of a successful fructose industry using 
raw fructosans from barley is slight* The only assurance 
that may be offered is that the actual yield of fructosans 
from barley grown under normal field conditions will be 
extremely variable and difficult to predict. 

Attempts to breed a high-fructosan barley would be 
premature in the face of present knowledge of the large and 
profound effect of the many factors, both environmental and 
physiological, on fructosan storage. 

Some indication of what remains to be done before 
our knowledge of fructosan formation is complete is revealed 
in the following quotation taken from Archbold (4): "Only 
when it can be stated with certainty under which circumstances 
glucose and fructose are interconvertible in plants, and when 
some knowledge has been acquired of the mechanisms involved 
both in such conversions and in polymerisation, will real 
progress in our views on fructosan formation be possible." 

In the opinion of the writer, more information and 
precise results from this study would be obtained if some of 
the following modifications were adopted: 

1. Samples for analysis should be based on a known 
number of plants and results reported on the quantity of sugar 
and dry matter per plant. 

2. Since the variation in sugar content with time is 
great, analysis should be made more frequently than once a 
week. 


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66 - 


3. Dry matter content and sugar concentrations based 
on the percentage of green or fresh weight are difficult to 
estimate with any degree of precision if free water adhering 
to plants is not rapidly and completely removed. At present 
no means of removing such water completely are known, but the 
use of blotters has been suggested. 

4. The period selected in this investigation for esti¬ 
mations of "overnight" changes in sugar, water, and dry 
matter was very unsatisfactory as varying hours of sunlight 
were included. Plants for the estimations of the effect of 
daylight should be selected near sun-down. Plants selected 
for the "morning” collection should be covered the evening 
previous or collected soon after sunrise. 

Finally, the following points should be borne in 
mind in any future work on the problem of carbohydrate 
storage in plants: 

1. The techniques for analysis of sugar in plants, now 
generally in use, while greatly improved over those formerly 
employed, require further refinement. More information as to 
sources of the glucose that are at present undefined is 
required before a completely satisfactory technique for esti¬ 
mation of sucrose and fructosan can be devised. 

2. The large losses of sugars observed during ripening 
or drying of plants require explanation. The suggestion that 


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67 


these and overnight losses are related to respiration has 
been advanced but is by no means conclusive, 

3. Recent observations made by Archbold and Datta (7), 

Gregory (18), and Richards (16) on carbohydrate storage and 
utilization indicate that nitrogen compounds are involved. 

The study of the interrelations of these two groups of com¬ 
pounds appears timely, 

ACKNOWLEDGEMENTS 


The writer’s thanks and appreciation are extended 
to the supervisor of this project, Dr. A. G. McCalla, for 
helpful criticism and advice during the progress of the 
investigation. A. E. Harper assisted in the routine analysis 
and G. M, Tosh in the preparation of equipment and photo¬ 
graphs. 

Financial assistance was supplied by the National 


Research Council, Ottawa 



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REFERENCES 


ARCHBOLD, H.K. Ripening processes in the apple and the 
relation of time of gathering in the chemical 
changes in cold storage, Ann. Bot. 46:407-459. 1932 

___ Physiological studies in plant nutrition: 

The role of fructosans in the carbohydrate metabolism 
of the barley plant. Materials used and method of 
sugar analysis employed. Ann. Bot. N.S. 2:183-202. 
1938. 


__Physiological studies in plant nutrition: 

The role of fructosans in the carbohydrate metabolism 
in the barley plant. Ann. Bot, N.S. 2:403-435. 1938 

Fructosans in the monocotyledons: A review 
New Phytologist 39:185-219. 1940. 

___ Physiological studies in plant nutrition: 

~ Experiments with barley on defoliation and shading of 
the ear in relation to sugar metabolism. Ann. Bot. 
N.S. 6:487-531. 1942. 

and BARTER, A.M. A fructose anhydride from 
the leaves of the barley plant. Biochem. Jour. 29: 
2689. 1935. 

and DATTA, C. Sugar metabolism in the bar- 
ley stem"in relation to ear development. Ann. Bot. 
N.S. 8(32):363-384. 1944. 

_ ^ and MUKERJEE, B.N. Physiological studies 

in plant nutrition: Carbohydrate changes in the 
several organs of the barley plant during growth, 
with special reference to the development and ripen¬ 
ing of the ear. Ann. Bot. N.S. 6:1-41, 1942. 

BARNELL, H.R. Seasonal changes in the carbohydrates of 
the wheat plant. New Phytologist 35:229-266. 1936. 

Distribution of the carbohydrates between 
component parts of the wheat plant at various times 
of the season. New Phytologist 37:85-112. 1938. 

CURTIS, O.T. The food content of forage crops as in¬ 
fluenced by the time of day at which they are cut. 
Jour. Am. Soc. Agron. 36:401. 1944. 













' * o 

loohiedo afiit at \ i.clLovoobo tv to n..cl$rd,3*i 

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■ 11 : .. a t . il , o 

... r : ; ' : . : ... To 

* ■ . ' ' - : ' 

I ft 0 . . 

■\ . . 70 ,...d 0 ^...r \xi£c.o$ Oi.:Tl 

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» 









69 


12. GREGORY, F.G. Mineral nutrition of plants. Ann. Rev. 

Biochem. 6:577. 1937. 

13. HAWORTH, W.N., HIRST, E.L., and LYME, R.R. A water- 

soluble polysaccharide from barley leaves. Biochem. 
lour. 31:786. 1937. 

14. NORMAN, G.A. The composition of forage crops. Biochem. 

lour. 30:1354-1362. 1936. 

15. REIGH, W.S. Separation of glucose and fructose. 

Biochem, lour. 33:1000. 1939. 

16. RICHARDS, F.I. Physiological studies in plant nutrition: 

The relation of respiration rate to the carbohydrate 
and nitrogen metabolism of the barley leaf as deter¬ 
mined by phosphorus and potassium supply. Ann. 

Bot. N.S. 2:491-534. 1938. 

17. RUSSELL, R.S. The effect of mineral deficiency on the 

fructosan metabolism of the barley plant, Ann. Bot. 
N.S. 2:865-882. 1938. 

18. SHAFFER, S0M0GYI. Gopper-iodometric reagents for sugar 

determination. lour. Biol. Chem. 100:695. 1933. 

19. VAN DER PLANK, I.E. The estimation of sugars in the leaf 

of the mangold, Biochem. Jour. 30:457-483. 1936. 

20. WILLARD, G.J. Afternoon vs. morning cutting of alfalfa. 

(Also notes by WOODWARD, T.E., SHEPHERD, J.B., 
TYSDALE, W.H. and CURTIS, O.T.). Jour. Am. Soc. 
Agron. 36:937-952. 1944. 

21. YEMM, E.W. Methods for determination of carbohydrates 

in leaves, Proc. Royal Soc. of London, Series B. 
117:483-504. 1935. 


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